title: Testing promotes long-term learning via stabilizing activation patterns in a large network of brain areas.
abstract: The testing effect refers to the phenomenon that repeated retrieval of memories promotes better long-term retention than repeated study. To investigate the neural correlates of the testing effect, we used event-related functional magnetic resonance imaging methods while participants performed a cued recall task. Prior to the neuroimaging experiment, participants learned Swahili-German word pairs, then half of the word pairs were repeatedly studied, whereas the other half were repeatedly tested. For half of the participants, the neuroimaging experiment was performed immediately after the learning phase; a 1-week retention interval was inserted for the other half of the participants. We found that a large network of areas identified in a separate 2-back functional localizer scan were active during the final recall of the word pair associations. Importantly, the learning strategy (retest or restudy) of the word pairs determined the manner in which the retention interval affected the activations within this network. Recall of previously restudied memories was accompanied by reduced activation within this network at long retention intervals, but no reduction was observed for previously retested memories. We suggest that retrieval promotes learning via stabilizing cue-related activation patterns in a network of areas usually associated with cognitive and attentional control functions.
authors.affiliation: Department of Cognitive Science, Budapest University of Technology and Economics, Budapest, Hungary.
authors.forename: Attila
authors.initials: A
authors.lastname: Keresztes
authors.affiliation: Institute of Psychology, University of Regensburg, Regensburg, Germany.
authors.forename: Daniel
authors.initials: D
authors.lastname: Kaiser
authors.affiliation: Department of Cognitive Science, Budapest University of Technology and Economics, Budapest, Hungary, Institute of Psychology, University of Regensburg, Regensburg, Germany, DFG Research Unit Person Perception, Friedrich-Schiller-University of Jena, Jena, Germany and Institute of Psychology, Friedrich-Schiller-University of Jena, Jena, Germany.
authors.forename: Gyula
authors.initials: G
authors.lastname: Kovács
authors.affiliation: Department of Cognitive Science, Budapest University of Technology and Economics, Budapest, Hungary.
authors.forename: Mihály
authors.initials: M
authors.lastname: Racsmány
mesh_terms: D000328:Adult; D000704:Analysis of Variance; D001921:Brain / Q000098:blood supply / Q000502:physiology*; D001931:Brain Mapping; D005260:Female; D006801:Humans; D007091:Image Processing, Computer-Assisted; D008279:Magnetic Resonance Imaging; D008297:Male; D011939:Mental Recall / Q000502:physiology*; D009434:Neural Pathways / Q000098:blood supply / Q000502:physiology; D010100:Oxygen / Q000097:blood; D010153:Paired-Associate Learning / Q000502:physiology*; D011930:Reaction Time / Q000502:physiology; D013997:Time Factors; D014706:Verbal Learning; D055815:Young Adult
chemical_list: D010100:Oxygen
keywords: fMRI; forgetting; long-term learning; retrieval; testing effect